The Hypocreales of New Zealand. I

The Hypocreales ol JNew Zealand. I

By Joan M. Dingley, Plant Diseases Division,

Department of Scientific and Industrial Research

[Read before the Auckland Institute, February 15, 1950;
received by Editor , February 22, 1950 .]

The Hypocreales are fungi belonging to the Ascomycetes, as their
sexual spores are developed in an ascus. As in the Sphaeriales, the
asci are borne in a well-developed perithecium. The perithecial walls
are fleshy, often membranous, but not dark coloured, hard or brittle.
Paraphyses are absent from the ascogenous tissue, although pseudo-
paraphyses are present within the perithecium. For many years this
order has been separated from the Sphaeriales by its fleshy perithecia.
Recent work by Miller (1949) based on developmental studies of
Sphaerostilbe aurantiicola (Berk and Br.) Petch by Luttrell (1944)
has shown that tissue formed within the perithecium prior to the
development of the asci become, in the mature perithecium, evanescent,
but remain as vertical or irregularly developed filaments often arising
from the apex of the perithecium. Seeler (1940) in discussing the
genus Thyronectria stated that paraphyses were lacking, but that
evanescent branching filaments were sometimes present. He observed
that these filaments differ from paraphyses as they grow downwards
from the position later occupied by the ostiole and fill the perithecial
cavity with a gelatinous substance. These observations agree with those
published by Luttrell. Examination of New Zealand material has
shown that paraphyses were absent, but evanescent tissue within
the perithecium was present in some species, even in those genera,
e.g. Claviceps and Cordyceps, that Nannfeldt (1932) included in the
family Clavicipitaceae. In this present account this family has been
retained in the Hypocreales, not in the Sphaeriales, the position
suggested for it by Miller.

Seaver (1909), in his treatment of the North American Hypo¬
creales, and Petch (1938) the British Hypocreales, divided the order
into two families, Nectriaceae and Hypocreaceae, on stromatic and
perithecial characters. Spore characters were retained by both for
generic and specific separation. Throughout the New Zealand species
studied, stromatic characters varied with the substratum, therefore
this separation between the families Hypocreaceae and Nectriaceae
was dropped. Ascal characters as defined by Nannfeldt were used
to separate the families Hypocreaceae and Clavicipitaceae. Spore
characters were used for generic and specific separation.

Morphology

Perithecia and stroma are superficially arranged on the host sub¬
stratum. When the stroma is absent, perithecia are scattered freely
on the host tissue, but if a stroma is present, the perithecia are eaespi-
tose or gregariously arranged, sometimes completely immersed in
the stromal tissues. The stroma is pulvinate, sometimes erumpent, or
effuse.

Transactions of the Royal Society of New Zealand
Yol. 79, Part 1, pp. 55-61, Plate 10

55

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Transactions

Throughout the order perithecia possess a well-defined wall, usually
pseudoparenchymatous, but in some species the mycelial walls are
densely thickened and pigmented and the nature of the tissue of the
perithecial wall is difficult to discern. In other species pigmented
inclusions within the cells gave colour to the perithecium. Where
perithecia are immersed in the stromal tissues, the perithecial wall is
poorly developed, but it is always distinct. An ostiolum is always
present, usually papillate and lined with periphyses. The stroma
varies from a loosely aggregated mat of mycelium to a well-developed
pulvinate pseudoparenchymatous structure.

Asci develop from ascogenous hyphae at the base of the peri¬
thecium. They typically contain eight spores, although in some genera,
the spores fragment within the ascus to form numerous part-spores.
Within the order asci are divided into two types according to the
method in which the spores are liberated.

(a) No special mechanism is associated with the liberation of spores.
The wall at the apex of the ascus ruptures and mature spores
are freed. Asci are thin walled, clavate, cylindrical or elliptical,
typical of asci of the family Hypoereaceae.

(b) The contents of the mature ascus force off a definite cap or
apical segment similar to the operculum in asci of the Pezizales.
Asci are cylindrical and thick walled, truncate and thickened
at the apex. This condition is characteristic of the family
Clavicipitaceae.

Paraphyses are absent; it is possible that earlier records of para-
physes were based upon misidentification of pseudoparaphyses or
young asci.

Spores may be non-septate, uniseptate, multiseptate or muriform.
In some genera the spores fragment at the septa to form unicellular
part spores. They are typically hyaline, although lightly pigmented
spores are characteristic of some species.

In New Zealand few groups of the Ascomycetes have been studied.
In Hooker’s Handbook to the New Zealand Flora, published in 1867,
two species of Nectria, one Hypocrea and two species of Cordyceps
were listed and briefly described. Cooke (1879) listed and described
a few further collections sent to Kew from New Zealand. Cunningham
(1921) described species in the genus Cordyceps.

The present treatment is by no means complete, as collecting has
not been sufficiently extensive throughout the Dominion.

Family I. HYPOCREACEAE

Perithecia light or brightly coloured, walls pseudoparenchymatous,
fleshy or membraneous, rarely dark coloured, never hard or brittle;
perithecia scattered or aggregated on or immersed in a stroma. Asci
not operculate, spores liberated by rupturing of apical wall of ascus.
Paraphyses absent.

Key to Genera

Ascospores one-septate

Spores elliptical or fusiform, apiculate . .. .. 1. Hypomyceg

Spores oval, elliptical or fusiform, not apiculate . 2. Nectria

Ascospores one septate fragmenting into unicellular part-
spores within the ascus ,. . t ,, .. ., ,. 3. Hypocrea

Dingley— The Hypocreales of New Zealand

57

Ascospores multiseptate.

Spores with two or more transverse septa.

Perithecia red or lightly coloured .. .. .. 4. Calonectria

Peritliecia violet or black .. .. .. .. 5. Ghbberella

Spores muriform .. .. . . .. 6. Thyronectria

Hypomyces includes species with fleshy, often brightly coloured
perithecia, superficial or semi-immersed in an effuse subiculum. Spores
are one-septate, fusiform, apiculate and coarsely verrucose. Most
species parasitize other fungi. Although originally defined to include
only species with one-septate fusiform, apiculate spores, the genus
for a time was treated as a biological one to include all Nectria- like
fungi which parasitize other fungous fructifications. To-day the genus
is limited to its original definition. Sydow (1920) separated species
with unequally divided spores into the genus Apiocre a, but this
character seems too variable to justify this treatment, consequently
all species are included in Hypomyces. Species with smooth, elliptical
or oval one-septate spores are placed under Nectria.

Hypomyces Tulasne. Annales des Science Naturelles, ser. iv, vol. 13,

p. 11, 1860.

Clintoniella (Sace ) Rehm. Hedurigia, vol. 39, p. 223, 1900 (in part) ;

Apiocrca Sydow. Annales Mycologici , vol. 18, p. 186, 1920.

Perithecia superficial or immersed in an effuse, byssoid stroma;
perithecial wall pseudoparenchymatous, pigmented and hyaline. Asci
containing eight spores, paraphyses absent. Spores one-septate, fusi¬
form, apiculate, verrucose, hyaline or lightly pigmented.

Type Species. Hypomyces lactifluorum Schw.

Distribution. World wide.

Key to Species

Spores under 30/t in length.

Spores equally divided by septa.

Perithecia globose; spores 17-26 X 4-6/1
Perithecia pyriform; spores 26-30 X 4 -Gy

Spores unequally divided by septa
Spores over 30/t in length.

Perithecia superficial; spores 27-45 X 4-6/t ..

Perithecia immersed in byssoid stroma; spores
27-38 X 5-7g .

1. H. aurantius Tul.

2. H. roaellus (Alb. & Schw.)

Tul.

3. H. ohryso8permu8 Tul.

4. H. novae-zealandiae

Dingley

5. H. armeniacus Tul.

1. Hypomyces aurantius (Persoon) Tulasne. Selecta Fungorum
Carpologia, vol. 3, p. 48, 1865. (Plate 10, Pig. 3.)

Sphaeria aurantia Pers Synopsis Fung., p. 18, 1801; S. aurantia Pers.
ex Fr. Syst. Myc., vol. 2, p. 440, 1823; Nectria aurantia Fr. Sunvnta
Veg. Scand., p. 388, 1849.

Subiculum effuse, spreading over pileus of host, orange and umber,
margins floccose, light yellow or white, pigment granules present on
outer wall of mycelium; perithecia gregarious, crowded, superficial
or semi-immersed in subiculum, globose or oval, 0*2-0*3 X 0*4 mm.,
orange or ferrugineous, translucent, ostiole papillated; perithecial
wall pseudoparenchymatous, 50/* thick, cells 8-15 X 12-25/*, thin
walled, lightly pigmented, pigmented granules present among the cells.
Asci cylindrical, ends truncate, thin-walled, 80-140 x 4-6/*, 8 spored,
spores obliquely uniseriate. Spores one-septate, equally divided into
two cells, fusiform, slightly curved, apiculate, 17-26 X 3-6/* hyaline
and verrucose.

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Transactions

Type Locality: Europe.

Distribution: Europe, North and South America, West Indies,
Australia, New Zealand.

Habitat: On fructifications of Polyporaceae. Coriolus velutinus
(Pers. ex Fr.) Quel. Auckland, Mount Albert, September, 1948,
D. W. McKenzie. Fomitopsis hemitephrus (Berk.) G. H. Cunn. Otago,
Otautau, December, 1946, G. B. Rawlings (2 col.). Irpex brevis Berk.
Auckland, Waipoua, September, 1949, Joan M. Dingley. Poria sp.
Auckland, Clevedon, August, 1948, Joan M. Dingley.

A distinct species, easily recognised by its bright orange, effuse
subiculum which covers the pileus of the host. Perithecia are darker
coloured hut translucent. Colenso (1886) first recorded this species for
New Zealand from material collected in Hawke’s Bay and determined
at Kew by M. C. Cooke.

No conidial stage is present, but Petch (1938) described Diplo-
cladium penicillioides Sacc. as its conidial form. The spores are borne
on a. stalked head or phialides, conidia are one-septate, 9-16 X 6-9/a,
hyaline. He also described chlamydospores 18-48 X 12-18/a, one- to
three-septate, reddish-brown.

2. Hypomyces rosellus (Alb. and Schw.) Tulasne. Annales des Science
Naturelles, ser. iv, vol. 13, p. 12, 1860. (Plate 10, Fig. 6.)

Sphaeria rosella Alb. and Schw. Conspect. Fung p. 35, 1805; 8. rosella
Alb. and Sch. ex Fr. Syst. Myc ., vol. 2, p. 441, 1822.

Subiculum effuse, white or rose coloured, mycelium compacted into
a pseudoparenehymatous layer 0 3 mm. thick, hyphae thin-walled and
lightly pigmented. Perithecia gregarious, superficially arranged, pyri¬
form, 0 2—0-25 X 0 3 mm., collapsing when dry, ostiole papillated,
hairy, rose or vinaceous coloured; perithecial wall pseudoparen¬
ehymatous, 40/a thick, cells 8-14 p diameter, rectangular or cuboid, cell
walls pigmented. Asci cylindrical, ends truncate, thin walled, 90-200
X 4-6/a, 8 spored, uniseriate, sometimes obliquely arranged. Spores
unisept ate, equally divided into two cells, fusiform or elliptical, apicu-
late, occasionally allantoid, 26—30 X 4-6/a, hyaline or verrucose.
Conidial stage.- Conidia elliptical or obtuse, sometimes allantoid,
I-3-septate, 16-28 X 7-10/a, smooth, hyaline, sometimes constricted
at septa. Conidiophores verticilloid or irregularly arranged on upright
hypha Dactylium dendroides Fr., Syst. Myc., vol. 3, p. 413, 1829.

Type Locabty: Germany.

Distribution: North and South America, West Indies, Australia.
New Zealand.

Habitat: On decaying fungus fructifications. Poly poms sp.
Auckland, Waitakere Ra., off Anawhata Rd., 1,000 ft., August, 1948,
Joan M. Dingley. Unknown host. Auckland, Otau, May, 1949, Joan M.
Dingley; Wellington, Weraroa, August, 1919, G. H. Cunningham.

Seaver (1910) described two conidial forms, a species of Tricho-
thecium as well as a Dactylium sp.

The. Wellington collection was determined by C. G. Lloyd as
Clintoniella rosella (Alb and Schw.), Clintoniella being erected by
Saccardo (1883) as a sub-genus of Hypocrea to include forms with
one-septate spores and an effuse stroma. According to Hoehnel (1918)
Rehm raised Saccardo’s sub-genus to generic rank, citing H. apiculata

Plate 10

Trans. Royal Society of N.Z., Vol. 79

To face page 58 ]

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Dingley —The Hypocreales of New Zealand

Peck as the type species. At this time some workers still regarded
Hypomyces as a biological genus and included in the genus Clintoniella
species morphologically similar to Hypomyces, but which did not
appear to parasitize other fungi. Seaver (1910) examined the co-type
of Peck’s type species, Hypocrea apiculata, and found it to be typical
of the genus Hypomyces and listed Rehm’s genus Clintoniella as a
synonym. Hoehne] {ibid.) examined species within Saccardo’s sub¬
genus and found them to belong to different genera within the
Hypocreaceae.

3. Hypomyces chrysospemms Tulasne. Annales des Sciences Natur -

elles, ser. iv, vol. 13, p. 16,1860.

Apiocrea ehrysosperma (Till ) Sydow. Annales Mycologici , vol. 18, p. 187,
1920.

Subiculum effuse, fioccose, white, mycelium thin walled, hyaline.
Perithecia gregarious, semi-immersed or superficial, globose, or oval,
0*2-0 -3 mm. diameter, light yellow, brown when dry, ostiole papil-
lated; perithecial wall pseudoparenchymatous, 20-25/a thick, cells
5-7 /Ji diameter, rectangular or cuboid, thin-walled and lightly pig¬
mented. Asci cylindrical, thin-walled, apex truncate, 90-120 X 4-6/a,
8-spored, obliquely uniseriate. Spores one-septate, cells unequal,
elliptical, fusiform and apiculate, 14-24 X4— 6/a, verrucose and hyaline.
Chlamydospores globose, thick-walled, 16-20 /a, tubereulate and yellow.
Sepedonium chrysospermum (Bull.) Fr. Syst. Myc., vol. 3, p. 438,
1829.

Type Locality: France.

Distribution: Europe, North America, Australia, New Zealand.

Habitat: Parasitic on fructifications of Boletus sp. Auckland, Wai-
takere Ra., Titirangi, June, 1946, Myra W. Carter; Swanson, May,
1948, Joan M. Dingley; Little Barrier Island, November, 1947,
Joan M. Dingley.

The bright yellow colour of the chlamydospores give the immature
subiculum a characteristic colour. Perithecia are rarely mature before
the pileus of the host has collapsed. Although not all spores are un¬
evenly divided by the septa all microscopic preparations contain both
evenly and unevenly divided spores. Petch (1938) described a conidial
stage as follows: “white effuse conidia, pyriform or oblong oval,
slightly constricted in the middle, continuous becoming one or two
septate 10 X 5/a to 30 X 12/a; conidiophore irregularly verticillioid.
often clustered.”

4. Hypomyces novae-zealandiae n.sp.* (Plate 10, Figs. 2, 5.)

Subiculum effusum, album, byssoidum. Perithecia gregaria in

subiculo, superficialia, globosa, vel pyriforma, 0*3-0 *5 mm. lutea,
ostiolum papillatum. Asci cylindrici, truncati 70-200 X 4-6/a. Sporae
uni-septatae, fusiformae, apiculatae 27-45 X 4-6/a, hyalinae verruco-
saeque.

Subiculum effuse, white, pale yellow, byssoid, mycelium pale
coloured. Perithecia gregarious, superficial, rarely semi-immersed in
stroma, globose sometimes pyriform, 0 3-0*5 mm., luteous or sulphur
yellow, translucent when fresh, ostiole papillated; perithecial wall

I arn indebted to Marjorie Newhook for the Latin translation.

60

Transactions

pseudoparenchymatous 30-40//, thick, cells rectangular or cuboid 5-1 Op
diameter, thin-walled, lightly pigmented yellow. Asci cylindrical, thin-
walled, truncate 70-200 X 4-6^, 8-spored, uniseriate. Spores one-
septate, equally divided into two cells, fusiform, apieulate 27-45 X
4 -6/a, hyaline and verrueose.

Type Locality: Waitakere Range, Auckland.

Disinbuhon: New Zealand.

Habitat: On pileus of Polyporus sp. Auckland, Waitakere Ra., off
Anawhata Rd., August, 1947, Joan M. Dingley. Type collection.

This species is separated from H. armeniacus Tul. by the superfici¬
ally arranged yellow perithecia and large fusiform spores. In growth
form it is similar to H. rosellus , but spores are much larger and
perithecia yellow, instead of red.

5. Hypomyces armeniacus Tulasne L. R. Annales des Science
Naturelles, ser. iv, vol. 13, p. 12, 1860. (Plate 10, Figs. 1 and 4.)
Hypomyces ochraceus (Peis ) Tul., Sel. Fung. Carp., vol. 3, p. 61, 1865;

Hypocrea apiculata Peck, 29th Hep. N.Y. State Mus ., p. 75, 1878;

Hypomyces terrestus Plowr. and Boud., Grevillea , vol. 8, p. 105, 1880;

Hypomyces apiculatus (Peck) Seaver, Mycologia, vol. 2, p. 73,’ 1910;

H. macrosporus Seaver, Mycoloyia, vol. 2, p. 80, 1910

Subiculum effuse, completely obliterating gills of host pileus, pale
flesh coloured, or ochraceous, mycelium compacted, hyaline. Perithecia
gregarious, completely immersed in subiculum, globose or oval 0-3-0 *4
X 0 4-0.5 mm., ochraceous or flesh coloured, ostiole papillated, trans¬
lucent, peritheeial wall pseudoparenchymatous 20//. thick, cells thin-
walled, lightly pigmented Asci cylindrical 120-160 X 5-8//,, 8-spored,
uniseriate, sometimes obliquely arranged. Spores one-septate, fusiform
with ends apieulate 25-38 X 5-7//,, verrueose, sometimes constricted at
septa, pale yellow.

Type Locality: France.

Distribution: Europe, North America, New Zealand.

Habitat: On pileus of Agarics. Lactarius sp. Wellington, Levin,
October, 1919, E. H. Atkinson.

Petch (1938) described a conidial stage for this species, Verti -
cillium agaricinnm Corda, with conidia obovate 11-21 X 9-12/* or
globose 10-13//.; he also recorded chlamydospores Blastotrichum
puccinioides Preuss, as compound 2-4 septate with large central cells
70-140 X 24-33//. reddish-purple and smooth. These stages were not
observed in the New Zealand collections.

Confusion exists as to the correct name of this species. Maire (1911)
concluded that after examining Tulasne’s type collection of H. armeni¬
acus this species agreed with Seaver’s H. macrosporous. Seaver (1910)
noted that his species H. macrosporous was probably H. ochraceus
(Pers.) Tul., but Persoon’s material was not available for comparison.
Maire (ibid.) stated that Tulasne in Selecta Fungorum Carpologia
(1865) changed the name of H. armeniacus to H. ochraceus presum¬
ably to identify the species with Sphaeria ochracea Pers., although
Tulasne himself was doubtful that these species were synonymous.
Fries (1822) listed Sphaeria ochracea as a variety of S. citrina
(Hypocrea citrina Fr.). Petch (1937) neglected Maire’s work and
referred this species to Hypomyces ochraceus (Pers.) Tul., listing

61

Dingley —The Hypocreales of New Zealand

Sphaeria ochracea Pers. as a synonym. H. terrestris Plow, and Bond,
was, according to Petch, erected because the authors did not realise
that the perithecial stage of this fungus seldom developed until t e
agaric had completely disintegrated and concluded that the species
was always found parasitic on the agaric and never terrestrial, in
other respects these species are identical. Petch also stated that
Miss Wakefield of Kew had determined that H. aptculatus (Feckj
Seaver was identical with Plowright’s species.

Literature Cited

Cooke, M. C , 1879. Grevillea, vol. 8, pp. 54-60.

Colenso, W., 1886. Trans. N.Z . Inst., vol. 19, pp. 301-313.

Cunningham, G H., 1921. Trans. N Z. Inst., vol. 53, pp. 372-382.

Hoehnel, F., 1918. Annales Mycologia, vol. 16, pp. 35-174.

Luttrell, E. S , 1944 Bull, of the Torrey Bot. Club, vol. 71, pp. 599-619.

Maire, Rene, 1911. Annales Mycoloyici, vol 9, pp. 315-325.

Miller, J. H., 1949. Mycologia, vol. 41, pp. 99—127.

Nannfeldt, J. A., 1932. Studien uber die Morphologie und Systcmatik der Nicht ■
Lickeniscerten Inoperculaten discomyceten, 368 pp.

Fetch, T., 1937. Journal of Botany, vol. 75, pp. 218-231.

__ U>38. Trans. British Mycological Soc ., vol 21, pp.

Saccardo, P. A, 1883. 8ylloge Fungorum, vol. 2, pp 1-958.

Seaver, E. J., 1909. Mycologia, vol. 1, pp. 41-76.
-- 1910. Mycologia, vol. 2, pp. 48-92.

Heeler, E W., 1940. J. of Arnold Arboretum.
Sydow, P. and H., 1920. Annales Mycologia,

Fig. 1 — 3. armeniacus
Fig. 2 — 3. novae-zelandiae
Fig. 3— 3. aurantius

, vol. 21, pp. 429-460.
vol. 18, pp. 178—197.

Fig. 4 — H. armeniacus
Fig. 5 — 3. novae-zelandiae
Fig. 6 — 3. rosellus